A palaeoecological investigation of long-term stand-scale ecological dynamics in semi-open native pine woods : contributing to conservation management in east Glen Affric

This thesis investigates past structure and dynamics of native Caledonian pine woodland, representing part of the western fringes of the northern European boreal woodlands. The biogeographical extent and Holocene history of the Scottish pine woods are well studied, yet questions remain at finer scales. This thesis is concerned with two factors over the recent Holocene oceanic period; (i) the long-term ecology within the woods; the spatio-temporal dynamics, the canopy structure, and community composition and continuity; and (ii) the former extent of the woods, especially the temporal pattern of the inferred easterly contraction in woodland over recent history. The importance of these factors - to ecologists, challenged with understanding the theories of equilibrium and non-equilibrium processes in long-lived woodland communities - and to conservation managers - challenged, by policy directives, with implementing the restoration and expansion of native woodlands - is discussed. Equally the identification and quantification of ecological detail over ecologically relevant temporal and spatial scales is an important challenge for palaeoecology. The thesis therefore applies fine spatial resolution pollen analyses over a network of sites, within, and at the western edge of, the extant woodland zone in east Glen Affric. Correlation between these individual site histories develops a detailed view of the grain and extent of woodland within the landscape, previously missing from western pine woods. Careful attention is paid to the interpretive potential and limitations of fine resolution palaeoecology; especially with regard (i) to techniques that can identify and spatially quantify stand-scale structure and community composition via reference to modern analogues; and (ii) to defining chronologies and elucidating rates and patterns of temporal change. The opportunities and limitations of the technique are explored and discussed, to ensure an understanding of the rigour and potential of the palaeoecological contribution to ecological research and to provide an evidence base for conservation. The application of 210Pb dating using the CRS and CIC models is explored. The CRS model is confirmed as suitable for peat deposits, but its use may mask fluctuations in peat sedimentation rate, which may be illustrated by the CIC model. The value of multiple 14C assays for each core and a need for a new approach to chronologies for application to fine-scale palaeecological studies is discussed. The relevant source area around the small basins in this semi-open pinewood is tentatively confirmed at 20 m from the pollen source. Tentative pollen productivity estimates for five key taxa in this ecosystem are presented. The temporal stability of native woodland in Glen Affric is confirmed to the eastern part of the extant woodland zone; but challenged to the west, where the open and semi-open landscape has a long history. Woodland diversity decreased over the last c. 200 years, and past woodland also shows a greater ground flora diversity. The western extent of the Caledonian woodland in this landscape may have changed little in the last c. 4000 years. Fluctuations, some clearly cyclical, in heath, and in woodland, communities are identified in the pollen record. The former may be aligned to changes in grazing regime or climatic shifts; and the latter to autochthonous shifts important in the maintenance of suitable edaphic conditions for the continuity of woodland. Former woodland is confirmed as likely to have been open in structure and mixed in tree species composition. The results presented here suggest that some caution should be applied to use of the term Caledonian, or native ‘pine’ forest: ‘Caledonian forest’ may better reflect the heterogeneity of past forests, particularly the importance, and persistence, of birch. The implications for conservation management and restoration are discussed. It may be difficult to establish a sustainable woodland to the west of the extant stands, and any pine woodland here may need to be mixed with stands of broadleaved trees to maintain or restore soil structure and ecological function. The landscape to the west may have been open for several thousands of years, and consideration of this is required when managing for the future to prevent loss of biodiversity.EThOS - Electronic Theses Online ServiceUniversity of Stirling : Forest ResearchGBUnited Kingdo

Spatial patterns in soil organic matter dynamics are shaped by mycorrhizosphere interactions in a treeline forest

Aims In the Swedish sub-Arctic, mountain birch (Betula pubescens ssp. czerepanovii) forests mediate rapid soil C cycling relative to adjacent tundra heaths, but little is known about the role of individual trees within forests. Here we investigate the spatial extent over which trees influence soil processes. Methods We measured respiration, soil C stocks, root and mycorrhizal productivity and fungi:bacteria ratios at fine spatial scales along 3 m transects extending radially from mountain birch trees in a sub-Arctic ecotone forest. Root and mycorrhizal productivity was quantified using in-growth techniques and fungi:bacteria ratios were determined by qPCR. Results Neither respiration, nor root and mycorrhizal production, varied along transects. Fungi:bacteria ratios, soil organic C stocks and standing litter declined with increasing distance from trees. Conclusions As 3 m is half the average size of forest gaps, these findings suggest that forest soil environments are efficiently explored by roots and associated mycorrhizal networks of B. pubescens. Individual trees exert influence substantially away from their base, creating more uniform distributions of root, mycorrhizal and bacterial activity than expected. However, overall rates of soil C accumulation do vary with distance from trees, with potential implications for spatio-temporal soil organic matter dynamics and net ecosystem C sequestration

Draft Genome Sequence of Methylocella silvestris TVC, a Facultative Methanotroph Isolated from Permafrost

Permafrost environments play a crucial role in global carbon and methane cycling. We report here the draft genome sequence of Methylocella silvestris TVC, a new facultative methanotroph strain, isolated from the Siksik Creek catchment in the continuous permafrost zone of Inuvik (Northwest Territories, Canada)

Transitions in Arctic ecosystems: ecological implications of a changing hydrological regime

Numerous international scientific assessments and related articles have, during the last decade, described the observed and potential impacts of climate change as well as other related environmental stressors on Arctic ecosystems. There is increasing recognition that observed and projected changes in freshwater sources, fluxes, and storage will have profound implications for the physical, biogeochemical, biological and ecological processes and properties of Arctic terrestrial and freshwater ecosystems. However, a significant level of uncertainty remains in relation to forecasting the impacts of an intensified hydrological regime and related cryospheric change on ecosystem structure and function. As the terrestrial and freshwater ecology component of the Arctic Freshwater Synthesis we review these uncertainties and recommend enhanced coordinated circumpolar research and monitoring efforts to improve quantification and prediction of how an altered hydrological regime influences local, regional and circumpolar-level responses in terrestrial and freshwater systems. Specifically, we evaluate i) changes in ecosystem productivity; ii) alterations in ecosystem-level biogeochemical cycling and chemical transport; iii) altered landscapes, successional trajectories and creation of new habitats; iv) altered seasonality and phenological mismatches; and, v) gains or losses of species and associated trophic interactions. We emphasize the need for developing a process-based understanding of inter-ecosystem interactions, along with improved predictive models. We recommend enhanced use of the catchment-scale as an integrated unit of study, thereby more explicitly considering the physical, chemical and ecological processes and fluxes across a full freshwater continuum in a geographic region and spatial range of hydro-ecological units (e.g., stream-pond-lake-river-near shore marine environments)

Whole-crown 13C-pulse labelling in a sub-arctic woodland to target canopy-specific carbon fluxes

Climate change-driven increases in plant productivity have been observed at high northern latitudes. These trends are driven, in part, by the increasing abundance of tall shrub and tree species in arctic ecosystems, and the advance of treelines. Higher plant productivity may alter carbon (C) allocation and, hence, ecosystem C cycling and soil C sequestration. It is important to understand the contributions that the newly established canopy forming overstorey species makes to C cycling in these ecosystems. However, the presence of a dense understorey cover makes this challenging, with established partitioning approaches causing disturbance and potentially introducing measurement artefacts. Here, we develop an in situ whole-crown 13C-pulse labelling technique to isolate canopy C fluxes in areas of dense understorey cover. The crowns of five mountain birch (Betula pubescens ssp. czerepanovii) trees were provided with a 13CO2 pulse using portable field equipment, and leaf samples were collected from neighbouring con-specific trees and hetero-specific understorey shrubs on days 1–10 and 377 post-crown labelling. We found effective and long-term enrichment of foliage in labelled trees, but no evidence of the 13C-signal in con- or hetero-specific neighbouring trees or woody shrubs. This method is promising and provides a valuable tool to isolate the role of canopy tree species in ecosystems with dense understorey cover.Output Status: Forthcoming/Available Onlin

Shrub expansion in the Arctic may induce large-scale carbon losses due to changes in plant-soil interactions

Background Tall deciduous shrubs are increasing in range, size and cover across much of the Arctic, a process commonly assumed to increase carbon (C) storage. Major advances in remote sensing have increased our ability to monitor changes aboveground, improving quantification and understanding of arctic greening. However, the vast majority of C in the Arctic is stored in soils, where changes are more uncertain. Scope We present pilot data to argue that shrub expansion will cause changes in rhizosphere processes, including the development of new mycorrhizal associations that have the potential to promote soil C losses that substantially exceed C gains in plant biomass. However, current observations are limited in their spatial extent, and mechanistic understanding is still developing. Extending measurements across different regions and tundra types would greatly increase our ability to predict the biogeochemical consequences of arctic vegetation change, and we present a simple method that would allow such data to be collected. Conclusions Shrub expansion in the Arctic could promote substantial soil C losses that are unlikely to be offset by increases in plant biomass. However, confidence in this prediction is limited by a lack of information on how soil C stocks vary between contrasting Arctic vegetation communities; this needs to be addressed urgently

Rhizosphere allocation by canopy-forming species dominates soil CO2 efflux in a subarctic landscape

In arctic ecosystems, climate change has increased plant productivity. As arctic carbon (C) stocks are predominantly located below ground, the effects of greater plant productivity on soil C storage will significantly determine the net sink/source potential of these ecosystems, but vegetation controls on soil CO2 efflux remain poorly resolved. To identify the role of canopy‐forming species in below‐ground C dynamics, we conducted a girdling experiment with plots distributed across 1 km2 of treeline birch (Betula pubescens) forest and willow (Salix lapponum) patches in northern Sweden and quantified the contribution of canopy vegetation to soil CO2 fluxes and below‐ground productivity. Girdling birches reduced total soil CO2 efflux in the peak growing season by 53% ‐double the expected amount, given that trees contribute only half of the total leaf area in the forest. Root and mycorrhizal mycelial production also decreased substantially. At peak season, willow shrubs contributed 38% to soil CO2 efflux in their patches. Our findings indicate that C, recently fixed by trees and tall shrubs, makes a substantial contribution to soil respiration. It is critically important that these processes are taken into consideration in the context of a greening arctic since productivity and ecosystem C sequestration are not synonymous

Resistance of subarctic soil fungal and invertebrate communities to disruption of below-ground carbon supply

The supply of recent photosynthate from plants to soils is thought to be a critical mechanism regulating the activity and diversity of soil biota. In the Arctic, large-scale vegetation transitions are underway in response to warming, and there is an urgent need to understand how these changes affect soil biodiversity and function. We investigated how abundance and diversity of soil fungi and invertebrates responded to a reduction in fresh below-ground photosynthate supply in treeline birch and willow, achieved using stem girdling. We hypothesised that birch forest would support greater abundance of ectomycorrhizal (ECM) fungal species and fauna than willow shrubs, and that girdling would result in a rapid switch from ECM fungi to saprotrophs as canopy supply of C was cut, with a concomitant decline in soil fauna. Birch forest had greater fungal and faunal abundance with a large contribution of root-associated ascomycetes (ericoid mycorrhizal fungi and root endophytes) compared to willow shrub plots, which had a higher proportion of saprotrophs and, contrary to our expectations, ECM fungi. Broad-scale soil fungal and faunal functional group composition was not significantly changed by girdling, even in the third year of treatment. Within the ECM community, there were some changes, with genera that are believed to be particularly C-demanding declining in girdled plots. However, it was notable how most ECM fungi remained present after 3 years of isolation of the below-ground compartment from contemporary photosynthate supply. Synthesis. In a treeline/tundra ecosystem, distinct soil communities existed in contrasting vegetation patches within the landscape, but the structure of these communities was resistant to canopy disturbance and concomitant reduction of autotrophic C inputs

Tree planting in organic soils does not result in net carbon sequestration on decadal timescales

Tree planting is increasingly being proposed as a strategy to combat climate change through carbon (C) sequestration in tree biomass. However, total ecosystem C storage that includes soil organic C (SOC) must be considered to determine whether planting trees for climate change mitigation results in increased C storage. We show that planting two native tree species (Betula pubescens and Pinus sylvestris ), of widespread Eurasian distribution, onto heather (Calluna vulgaris ) moorland with podzolic and peaty podzolic soils in Scotland, did not lead to an increase in net ecosystem C stock 12 or 39 years after planting. Plots with trees had greater soil respiration and lower SOC in organic soil horizons than heather control plots. The decline in SOC cancelled out the increment in C stocks in tree biomass on decadal timescales. At all four experimental sites sampled, there was no net gain in ecosystem C stocks 12–39 years after afforestation—indeed we found a net ecosystem C loss in one of four sites with deciduous B. pubescens stands; no net gain in ecosystem C at three sites planted with B. pubescens ; and no net gain at additional stands of P. sylvestris . We hypothesize that altered mycorrhizal communities and autotrophic C inputs have led to positive ‘priming’ of soil organic matter, resulting in SOC loss, constraining the benefits of tree planting for ecosystem C sequestration. The results are of direct relevance to current policies, which promote tree planting on the assumption that this will increase net ecosystem C storage and contribute to climate change mitigation. Ecosystem‐level biogeochemistry and C fluxes must be better quantified and understood before we can be assured that large‐scale tree planting in regions with considerable pre‐existing SOC stocks will have the intended policy and climate change mitigation outcomes

Exploring drivers of litter decomposition in a greening Arctic: Results from a transplant experiment across a tree-line (Forthcoming)

Decomposition of plant litter is a key control over carbon (C) storage in the soil. The biochemistry of the litter being produced, the environment in which the decomposition is taking place, and the community composition and metabolism of the decomposer organisms exert a combined influence over decomposition rates. As deciduous shrubs and trees are expanding into tundra ecosystems as a result of regional climate warming, this change in vegetation represents a change in litter input to tundra soils and a change in the environment in which litter decomposes. To test the importance of litter biochemistry and environment in determining litter mass loss, we reciprocally transplanted litter between heath (Empetrum nigrum), shrub (Betula nana) and forest (Betula pubescens) at a sub-arctic tree-line in Sweden. As expansion of shrubs and trees promotes deeper snow, we also used a snow fence experiment in a tundra heath environment to understand the importance of snow depth, relative to other factors, in the decomposition of litter. Our results show that B. pubescens and B. nana leaf litter decomposed at faster rates than E. nigrum litter across all environments, while all litter species decomposed at faster rates in the forest and shrub environments than in the tundra heath. The effect of increased snow on decomposition was minimal, leading us to conclude that microbial activity over summer in the productive forest and shrub vegetation is driving increased mass loss compared to the heath. Using B. pubescens and E. nigrum litter, we demonstrate that degradation of carbohydrate-C is a significant driver of mass loss in the forest. This pathway was less prominent in the heath, which is consistent with observations that tundra soils typically have high concentrations of ‘labile’ C. This experiment suggests that further expansion of shrubs and trees may stimulate the loss of undecomposed carbohydrate-C in the tundra.REF Compliant by Deposit in Stirling's Repositor